1urn:lsid:arphahub.com:pub:f9b2e808-c883-5f47-b276-6d62129e4ff4urn:lsid:zoobank.org:pub:245B00E9-BFE5-4B4F-B76E-15C30BA74C02Biodiversity Data JournalBDJ1314-28361314-2828Pensoft Publishers10.3897/BDJ.5.e10967109674957urn:lsid:arphahub.com:pub:6338ef08-97ca-58d8-b3eb-b8ae28d80e0durn:lsid:zoobank.org:pub:A9BBF50A-0E56-4251-B536-4A4EF87E6D56http://tb.plazi.org/GgServer/summary/0B5CFC468A16FFA7FF9AA756FFC5FFE4Taxonomic PaperTachinidaeBiodiversity & ConservationSystematicsCosta RicaCentral America and the CaribbeanCentral AmericaAmericasFive new species of Vibrissina Rondani (Diptera: Tachinidae) from Area de Conservación Guanacaste in Northwestern Costa RicaFlemingAJajfleming604@gmail.com1WoodD. Monty1SmithM. Alexhttps://orcid.org/0000-0002-8650-25752DapkeyTanya3HallwachsWinnie3JanzenDaniel H.31Agriculture Agri-Food Canada, Ottawa, CanadaAgriculture Agri-Food CanadaOttawaCanada2Department of Integrative Biology and the Biodiversity Institute of Ontario, Guelph, CanadaDepartment of Integrative Biology and the Biodiversity Institute of OntarioGuelphCanada3University of Pennsylvania, Philadelphia, United States of AmericaUniversity of PennsylvaniaPhiladelphiaUnited States of America

Corresponding author: AJ Fleming (ajfleming604@gmail.com).

Academic editor: Daniel Whitmore

2017180720175e109672810201608072017AJ Fleming, D. Monty Wood, M. Alex Smith, Tanya Dapkey, Winnie Hallwachs, Daniel H. JanzenThis is an open access article distributed under the terms of the Creative Commons Attribution License 4.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.AbstractBackground

We describe five new species in the genus Vibrissina Rondani from Area de Conservación Guanacaste (ACG). All species were reared from wild-caught sawfly larvae (Hymenoptera: Symphyta: Argidae and Tenthredinidae). We provide a morphological description of each species together with information on life history, molecular data, and photographic documentation.

With more than 8,500 described species classified into more than 1,500 genera (Irwin et al. 2003, O'Hara 2016) the family Tachinidae is one of the most diverse of all the insect families. The Neotropical Region has a vast and largely unknown fauna of tachinids, with a majority of names belonging to the Mexican and Brazilian fauna deriving from work conducted by early researchers, and from a limited number of localities (Wood 1985). The last major work on Neotropical Tachinidae, conducted by Guimaraes (1971), catalogued some 2,864 species, a number much larger than that of any other faunal region. Based on what has recently been discovered in Costa Rica (Smith et al. 2007, Fleming et al. 2014a, Fleming et al. 2014b, Fleming et al. 2015c, Fleming et al. 2015a, Fleming et al. 2015b) and what is already present in museum collections, this number is undoubtedly just a small fraction of what actually exists in nature.

The present study is part of an effort to document the tachinid species living within the terrestrial 120,000 km2 Area de Conservación Guanacaste (http://www.acguanacaste.ac.cr), and provide names for any new species. A comparison of tachinids collected during the ACG inventory with those present in the national collection in the Museo Nacional de Costa Rica (formerly INBio) shows minimal overlap of species, suggesting that the tachinid fauna in other parts of the country is quite different from that of ACG and requires much additional study (Janzen et al. 2012, Bertrand et al. 2014, Brown et al. 2014, Fernandez-Triana et al. 2014, Phillips-Rodríguez et al. 2014, Fernandez-Triana et al. 2015, Sharkey et al. 2015).

The genus Vibrissina Rondani, 1861 (Exoristinae: Blondeliini) originally included Rondani (1861) two nominal species from the Palearctic, Vibrissinaturrita (Meigen, 1824) and Vibrissinadebilitata (Pandellé, 1896). Three Asian species were added by Shima (1983), but the genus as a whole remained unstudied until Wood (1985) revised the tribe Blondeliini and 7 other genera under Vibrissina, bringing the total number of valid included species to 28. Among the genera synonymized by Wood (1985), Spathimeigenia Townsend represented the largest proportion of the newly added species. Almost all members of the genus Vibrissina are parasitoids on sawfly larvae in the families Argidae, Diprionidae, and Tenthredinidae (Wood 1985).

Vibrissina possesses the following characters, common to the tribe Blondeliini: prosternum setose; first postsutural supraalar bristle shorter than first postsutural dorsocentral bristle; bend of vein M rounded, forming an obtuse angle; subapical scutellar bristles long, stout, and divergent; and veins R4+5 and M ending at or near wing tip (Wood 1985).

Wood (1985) made a distinction between New World and Old World species of Vibrissina, the Old World species being characterized in large part by a bare parafacial, a character which is rare in New World species. All ACG-reared species of Vibrissina are parasitoids of sawfly larvae in the families Diprionidae and Tenthredinidae. This host specificity as parasitoids of sawfly larvae indicate that the various species placed by Wood (1985) under Vibrissina may actually form a monophyletic assemblage.

In this paper we describe five new species of Vibrissina using morphology and CO1 (cytochrome c oxidase I) gene sequences or “DNA barcodes”, and provide additional information on host preference.

This inventory has reared more than 750,000 wild-caught caterpillars and sawfly larvae since 1978. All frequencies of parasitization reported here need to be considered against this background inventory (Janzen et al. 2009, Janzen and Hallwachs 2011, Fernandez-Triana et al. 2014). The scope of our treatment of the genus Vibrissina is limited to species found within ACG.

It should be noted that this inventory searches some vegetation types and vertical strata much more thoroughly than others. This bias is due to the methods employed for collecting of specimens, which rely solely on those animals within reach of the collectors, up to 3m above the ground. Recent comparisons of reared species of parasitoids with those collected in the same place with hand nets or Malaise traps demonstrate that, to date, the estimated 1,100 species of tachinid flies reared by the inventory represent less than half the species of caterpillar (and caterpillar-like larvae) parasitizing Tachinidae present in ACG. The largest unsampled habitat is the foliage of the canopy that is higher than approximately 3–4 m above the ground.

Species accounts presented in this paper are deliberately brief and only include basic descriptions of body morphology and coloration commonly used in the identification of Tachinidae. The descriptions are complemented with a series of color photos of every species, used to illustrate the morphological differences among them. The morphological terminology used follows Cumming and Wood (2009). All dissections and photography were carried out following the methods detailed in O’Hara (1983) and Fleming et al. (2014a). Measurements and examples of parts of the terminalia are illustrated in Fig. 1.

Acronyms for depositories

CNC Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Canada

NHMUK Natural History Museum, London, United Kingdom (formerly British Museum of Natural History)

NHMW Naturhistorisches Museum Wien, Vienna, Austria

USNM National Museum of Natural History, Washington, D.C., U.S.A. (formerly United States National Museum)

Voucher specimen management

The procedures surrounding the management of voucher specimens has been detailed in previous papers in this series (Fleming et al. 2014a, Fleming et al. 2014b, Fleming et al. 2015a, Fleming et al. 2015b, Fleming et al. 2015c). In brief, caterpillars reared from the ACG efforts receive a unique voucher code in the format yy–SRNP–xxxxx; any parasitoid emerging from a caterpillar receives the same voucher code, and if/when later dealt with as an individual, receives a second unique voucher code in the format DHJPARxxxxxxx. The collateral data for each voucher code are available at: http://janzen.bio.upenn.edu/caterpillars/database.lasso.

All DHJPARxxxxxxx-coded tachinids have had one leg removed and sent for DNA barcoding to the Biodiversity Institute of Ontario (BIO) and its Center for Biodiversity Genomics (CBG) in Guelph, Canada. All collateral data and successful barcodes are permanently and publicly deposited in the Barcode of Life Data System (BOLD, www.boldsystems.org) (Ratnasingham and Hebert 2007), and subsequently migrated to GenBank. Each barcoded specimen also receives an accession number from the Barcode of Life Data System (BOLD) and GenBank. Since the inventory is continually adding new specimens, these can be found by searching the genus Vibrissina in BOLD.

All inventoried specimens discussed herein were collected under Costa Rican government research permits issued to DHJ, and Tachinidae samples were exported under permit by DHJ from Costa Rica to their final depository in the CNC in Ottawa, Canada. Tachinid identifications for the inventory were done by DHJ in coordination with a) visual inspection by AJF and DMW, b) DNA barcoding by MAS and c) databasing/correlation with host caterpillars by DHJ and WH via the inventory itself.

The date of capture cited for each specimen is the date of eclosion of the fly and not the date of capture of the caterpillar. Eclosion date is much more representative of the time when that fly species is on the wing than is the time of capture of the parasitized caterpillar. The “collector” is the parataxonomist who found the caterpillar, rather than the person who later retrieved the newly eclosed fly and processed it by freezing, pinning, labelling and oven-drying. The holotypes and paratypes of the newly-described species are housed in the Diptera collection of the Canadian National Collection (CNC).

Interim names of undescribed host species

Names of undescribed host species follow a standardized, interim naming system used for taxonomic units considered as distinct species and identified by DNA barcodes. The interim names are given in the format "Eois Janzen52", where the species epithet is composed of the name of the taxonomist who identified the species and a number. This prevents confusion with already described species while maintaining traceability of each undescribed species within the ACG project.

Barcoding methods

We analyzed DNA barcodes (the 5’ region of the cytochrome c oxidase I (CO1) gene (Hebert et al. 2004)) for all specimens of ACG Vibrissina. Barcodes were amplified from total DNA extracts that had been prepared from single legs using a standard glass fiber protocol (Ivanova et al. 2006). This 658 bp region near the 5’ terminus of the CO1 gene was then generated using standard insect primers (LepF1–LepR1) following established protocols for production and quality control (see Smith et al. 2006, Smith et al. 2007, Smith et al. 2008).

Male. Head: vertex 1/4–1/3 of head width; 1–3 reclinate orbital bristles; anteriormost reclinate orbital bristle distinctly longer than uppermost frontal bristle; ocellar bristles well developed, long, and divergent; eye bare or at most with minute, inconspicuous hairs all but invisible except under certain angles of light; parafacial on most New World species haired on lower half or more, bare in few New World species (Wood 1985); parafrontal bearing hairs interspersed among frontal bristles; lower margin of face at level of vibrissa not visible in profile; facial ridge bristled; subvibrissal ridge short, usually with 3 or fewer bristles; anterior margin of postgena concave, usually without a genal dilation; genal groove extensive, often sparsely tomentose (suggesting Dexiini); postgena bearing few widely spaced bristles, the anteriormost usually larger and distinctly isolated from the rest; arista minutely pubescent, usually distinctly thickened on basal fourth or fifth. Thorax: prosternum setose; proepisternum bare; postpronotum with 3 bristles in a straight line or slightly curved row; katepisternum with 2 or 3 bristles, when 3 present then two anterior to suture and one posterior; lateral scutellar bristles shorter than subapical bristles, curved medially or straight, divergent, as long as subapical bristles. Legs: fore tibia with 1 posterior bristle; mid tibia with 1 anterodorsal bristle. Wings: vein R4+5 setose, bearing only 2–3 bristles dorsally at base. Abdomen: mid-dorsal depression on syntergite 1+2 (ST1+2) extending almost to hind margin; two median marginal bristles on T3 and often also on ST1+2; row of marginal bristles on T4 and T5; 1–3 pairs of median discal bristles on T3–T5 (5th tergite sometimes bearing a complete row of discal bristles). Female as male except: 2 pairs of proclinate orbital bristles present; T3 and T4 ventolaterally flattened, their edges either studded with spines or stout bristles.

Diagnosis

Our observations of New World species of Vibrissina confirm those made by Wood (1985). One of the main characteristics of the genus in the New World is that all species possess hairs on the lower half of the parafacial, usually separated from the lowest frontal bristle by a bare gap, a character trait which can be used to distinguish them from the rest of the Blondeliini, with the exception of Lixophagaretiniae (Coquillett) and Erynniopsisantennata (Rondani) (both restricted to California and Oregon), and all species of Cryptomeigenia Brauer & Bergenstamm, which also have hairs on the lower half of the parafacial, having 4 or more bristles on the subvibrissal ridge. Wood (1985) also cites Enrogaliamorigera Reinhard and Istochetaaldrichi (Mesnil) as having hairs on the parafacial; however, in the case of both these species the hairs are most abundant on the upper half, decreasing in size and density toward the lower half. Old World species of Vibrissina, which lack parafacial hairs, can be separated only with difficulty from species of Eucelatoria Townsend, and can be separated only by the lack of a genal dilation.

Distribution

Widespread throughout the Palearctic, and from Mexico to Brazil in the Neotropical Region (Wood 1985).

Ecology

Almost all members of Vibrissina are parasitic on the caterpillar-like larvae of sawflies in the families Argidae, Diprionidae, and Tenthredinidae (Wood 1985). The present data from ACG inventoried larvae support this.

Female (not pictured, due to lack of photographic quality specimens). Length: 5–6mm. As male, with the exception of the following characters: thorax: dorsum and scutellum entirely silver tomentose; thoracic vittae distinct both pre- and postsuturally. Abdomen: entirely black in ground color, ventrolaterally flattened; all tergites lacking discal bristles; median marginal bristles present on ST1+2 and T3; T4 and T5 each with a row of marginal bristles; mid-ventral portion of T3–T5 tergites with a row of strong, stout spines.

Diagnosis

Vibrissinaalbopicta can be differentiated from its congeners by the following combination of traits: parafacial and parafrontal silver; thoracic tomentum silver-gray on both dorsal and lateral surfaces; abdominal ground color almost entirely black with only traces of dark brown; silver tomentum extending to over 30% of T3 and T4, but absent on T5; ST1+2 and T3 with only one pair of median marginals, T4 and T5 with a complete row of marginals; discals on T3 and T4 underdeveloped in male, absent in female.

Female (Fig. 3d, e, f). Length: 5–6mm. As male, with the exception of the following characters: thorax: dorsum of thorax and scutellum almost entirely gold tinged with no scutellar discals; thoracic vittae distinct both pre- and postsuturally; ventral katepisternal bristle very reduced compared to the other two. Abdomen: black ground color over its entirety; brassy tomentum along anterior 1/2 of tergites T3, T4, and T5; ventrolaterally flattened; no visible discal bristles on any segment; 1 pair of reduced marginal bristles on ST1+2 and T3; T4 and T5 each with 1 row of marginal bristles; mid-ventral portion of T3–T5 with a row of strong stout spines.

Diagnosis

Vibrissinadanmartinisp. n. can be differentiated from its congeners by the combination of the following traits: parafacial and parafrontal gold with parafacial haired; outer thoracic vittae reduced presuturally, not extending to suture, postsuturally also weak; ST1+2, T3, and 50% of T4 bearing orange ground color lateroventrally; abdominal tomentum extending over 60% of tergites T3, T4, and T5, with tomentum broken up by a dorsocentral dark stripe; in females, abdominal tomentum brassy-gold; marginal bristles reduced on ST1+2 and T3, rows of marginal bristles on T4 and T5.

Etymology

Vibrissinadanmartinisp. n. is dedicated to Mr. Dan Martin, now of Washington, D.C., formerly of Chicago, Illinois, in recognition of his administrative and philanthropic support of the biodiversity development concept that gave birth in 1989 to the INBio national biodiversity inventory collections, now part of Museo Nacional de Costa Rica, and his decades of steadfast support for the protection of tropical biodiversity in general, and specifically that of Costa Rica.

Distribution

Costa Rica, ACG, Prov. Guanacaste, dry forest, between 280–295m.

Ecology

Hosts: reared 37 times from larvae of the sawfly Durgoamattogrossensis Malaise (Argidae), which feed on the leaves of Bauhiniaungulata L. (Fabaceae).

Vibrissinahallwachsorumsp. n. can be differentiated from its congeners by the combination of the following traits: parafacial and parafrontal gold; tergite 5 bearing all black ground color with silver tomentum over its entirety; and only 1 pair of discal bristles on T3.

Etymology

Vibrissinahallwachsorumsp. n. is dedicated to Robert and Marianne Hallwachs of Philadelphia, Pennsylvania in recognition of their seminal support in acquiring the buildings in which the INBio national biodiversity inventory collections have grown and thrived since 1989, and which are now donated to the Museo Nacional de Costa Rica.

Distribution

Costa Rica, ACG, Prov. Guanacaste, cloud forest, at 1090m.

Ecology

Hosts: reared once from a larva of the sawfly Waldheimiainterstitialis (Cameron) (Tenthredinidae) feeding on the leaves of Hameliapatens Jacq. (Rubiaceae).

Vibrissinarandycurtisisp. n. can be differentiated from its congeners by the combination of the following traits: parafacial and parafrontal all gold; ground color of T5 black in both sexes; abdominal tomentum covering over 30% of ST1+2, T3, and T4; females bearing only 1 pair of marginal bristles on segments ST1+2 and T3, and a single row of marginal bristles on T4.

Etymology

Vibrissinarandycurtisisp. n. is dedicated to Mr. Randy Curtis of Arlington, Virginia in recognition of his seminal support in acquiring the land on which the INBio national biodiversity inventory collections grew and thrived since their founding in 1989, and still do, now as part of Museo Nacional de Costa Rica.

Female (Fig. 8d, e, f). Length: 5–7mm. As male with the exception of the following characters: abdomen: ventrolaterally flattened, bearing only one pair of discal bristles per segment; 1 pair of marginal bristles on ST1+2 and T3; mid-ventral portion of T3–T5 with a row of strong stout spines.

Diagnosis

Vibrissinarandyjonesisp. n. can be differentiated from its congeners by the combination of the following traits: parafacial and parafrontal all gold; tergite 5 of strong orange ground color at apex, this trait present in both males and females.

Etymology

Vibrissinarandyjonesisp. n. is dedicated to Mr. Randy Jones of Poland, Ohio, in recognition of his seminal support in acquiring the buildings in which the INBio national biodiversity inventory collections grew and thrived since their founding in 1989, and still do, now as part of Museo Nacional de Costa Rica.

Distribution

Costa Rica, ACG, Prov. Guanacaste, dry forest, between 280–300m.

Ecology

Hosts: reared 100+ times from larvae of Durgoamattogrossensis (Argidae) feeding on the leaves of Bauhiniaungulata (Fabaceae).

Vibrissinarobertwellsisp. n. can be differentiated from its congeners by the combination of the following traits: parafacial silver and parafrontal gold; tergite 5 of strong orange ground color over more than 50% of tergite, a trait most prominent in females.

Etymology

Vibrissinarobertwellsisp. n. is dedicated to Mr. Robert Wells of San José, Costa Rica in recognition of his legal manoeuvering and contract management for INBio’s purchase of the lands on which the national biodiversity inventory collections have grown and thrived since 1989, and which have now been donated to the Museo Nacional de Costa Rica.

A neighbor-joining (NJ) (Saitou and Nei 1987) tree based on Kimura 2-parameter was used to visually demonstrate the variation present within and between each species in the DNA barcode locus and is presented in Fig. 12. The variation illustrated is based on the evolutionary distances computed using the Tamura-Nei (TN) method (Tamura and Nei 1993). The TN method was selected due to the lowest BIC (Bayesian Information Criterion) scores of the Maximum Likelihood fits of 24 different nucleotide substitution models run in MEGA6 (Tamura et al. 2013). Interested readers can consult the Barcode of Life Data System (BOLD) (Ratnasingham and Hebert 2007) for all information associated with each sequence (including GenBank accessions) derived from each individual specimen using the DOI dx.doi.org/10.5883/DS-ASVIBRI.

Acknowledgements

We gratefully acknowledge the unflagging support of the team of ACG parataxonomists (Janzen et al. 2009, Janzen & Hallwachs 2011) who found and reared the specimens used in this study, and the team of biodiversity managers who protect and manage the ACG forests that host these tachinids and their hosts. The study has been supported by U.S. National Science Foundation grants BSR 9024770 and DEB 9306296, 9400829, 9705072, 0072730, 0515699, and grants from the Wege Foundation, International Conservation Fund of Canada, Jessie B. Cox Charitable Trust, Blue Moon Fund, Guanacaste Dry Forest Conservation Fund, Area de Conservación Guanacaste, Permian Global and University of Pennsylvania (DHJ & WH). This study has been supported by the Government of Canada through its ongoing support of the Canadian National Collection, Genome Canada, the Biodiversity Institute of Ontario, and the Ontario Genomics Institute (2008–0GI–ICI–03) (MAS), and by a Discovery Grant from the Natural Sciences and Engineering Research Council of Canada (MAS).

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Neighbor-Joining displaying the inter- and intra-specific variation within the DNA barcode region for the 6 species of Vibrissina Rondani reared from ACG. Tip labels include the species name|specimen accession number|host species; the holotype of each species is indicated with an asterisk. Habitus photographs in lateral view are presented for the holotypes of the new species. An ACG-reared voucher specimen is presented for V.albopicta.